Cribrilinid Bryozoans from Pleistocene Mediterranean Deep-Waters, with the Description of New Species

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Journal of Paleontology, page 1 of 23 Copyright © The Author(s), 2020. Published by Cambridge University Press on behalf of The Paleontological Society. This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. 0022-3360/20/1937-2337 doi: 10.1017/jpa.2020.93

Cribrilinid bryozoans from Pleistocene Mediterranean deep-waters, with the description of new species

Antonietta Rosso,1,2 Emanuela Di Martino,3* and Andrew N. Ostrovsky4,5

1Dipartimento di Scienze Biologiche, Geologiche e Ambientali, University of Catania, Corso Italia 57, 95129, Catania, Italy 2CoNISMa (Consorzio Interuniversitario per le Scienze del Mare), Piazzale Flaminio 9, 00196, Roma, Italy 3Natural History Museum, University of Oslo, Blindern, P.O. Box 1172, Oslo 0318, Norway 4Department of Invertebrate Zoology, Faculty of Biology, Saint Petersburg State University, Universitetskaja nab. 7/9, 199034, Saint Petersburg, Russia 5Department of Palaeontology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, Althanstr. 14, 1090, Vienna, Austria

Abstract.—Cribrilinid bryozoans originating from Pleistocene deep-water sediments from two localities near Messina (Sicily, Italy)—Capo Milazzo (Gelasian) and Scoppo (Calabrian)—were examined. Five cribrilinid species were found, three in each locality and time interval, with only one species shared. Three species, Cribrilaria profunda n. sp., Glab- rilaria transversocarinata n. sp., and Figularia spectabilis n. sp., are new to science. Of the two remaining species, Figu- laria figularis was already known from local fossil associations, whereas Glabrilaria pedunculata, a present-day Mediterranean species, is recorded for the first time as a fossil. New combinations are suggested for two species previously assigned to Puellina, Cribrilaria saldanhai (Harmelin, 2001) n. comb. and Cribrilaria mikelae (Harmelin, 2006) n. comb. The diagnosis of the genus Figularia was amended to include an erect growth morphology in addition to the encrusting form, and the occurrence of ooecia formed by the distal kenozooid. Following a literature revision of all species currently assigned to Figularia, the new combinations Vitrimurella capitifera (Canu and Bassler, 1929) n. comb. and Hayamiellina quaylei (Powell, 1967a) n. comb. are suggested, and problematic species are listed and briefly discussed.

UUID: http://zoobank.org/b7b36152-bf7b-4e00-b6ec-2614b2a58f1b

Introduction of the numerous Cretaceous representatives (e.g., Taylor and McKinney, 2006; Rosso et al., 2018), as well as phylogenetic Cribrilinidae Hincks, 1879 is an extremely large family of chei- analyses. Genus and species identification are often based on lostome bryozoans including 127 genera and more than 700 subtle morphological characters, such as those associated with living and fossil species to date, accounting for ∼3% of total the zooidal orifice and the suboral bar (e.g., Harmelin, 1970, bryozoan diversity (Bock, 2020). First appearing ca. 100 Ma, 1978, 2001, 2006; Bishop and Househam, 1987), which require in the Cenomanian, Cribrilinidae underwent a peak of diversifi- scanning electron microscopy (SEM), still lacking in the cation during the Santonian, greatly contributing to the radiation descriptions of numerous taxa. In fossil material, identification of cheilostomes in the Late Cretaceous (Cheetham, 1971; of taxa is also jeopardized by taphonomic filters, with abrasion, Jablonski et al., 1997 and references therein). This family is corrosion, partial dissolution and recrystallization obliterating one of the most species-rich in the present-day Mediterranean fine diagnostic characters. This is particularly true for species (Rosso and Di Martino, 2016), as well as in other regions of introduced in old publications, normally including only brief the world (e.g., Gordon et al., 2019). Cribrilinids exhibit a descriptions and often lacking proper illustrations. Descriptions typical and distinctive costate frontal shield, but also high and revisions of fossil cribrilinids based on detailed illustrations morphological variability, including different types of hetero- are scarce in the modern literature, especially for specific morphs (avicularia, kenozooids, articulated and non-articulated stratigraphic intervals (Berning, 2006; Taylor and McKinney, spines, etc.) and ovicell structures. A future subdivision of Cri- 2006; Di Martino and Rosso, 2015). In this context, this paper brilinidae into several families or subfamilies is very likely. A aims to: (1) document cribrilinid associations from Pleistocene more accurate definition of certain genera will, however, require deep-water habitats of southern Italy; (2) illustrate fossil repre- a thorough re-examination of the original material, particularly sentatives of some established species; (3) describe three new species; (4) amend the diagnosis of the genus Figularia Jullien, 1886, and provide a comparative morphological analysis of spe- *Corresponding author cies currently assigned to this genus; and (5) propose new 1 Downloaded from https://www.cambridge.org/core. Universitetsbiblioteket i Oslo (UiO), on 18 Dec 2020 at 15:19:10, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/jpa.2020.93 2 Journal of Paleontology:1–23

combinations for two species of Puellina and two species of Figularia.

Geological setting

North-eastern Sicily is part of the north Sicily Chain, which, in this sector, includes the Kabilo-Calabride crystalline basement (Paleozoic rocks of different metamorphic grade) and its sedimentary cover (i.e., discontinuous upper Miocene sediments unconformably covered by Plio-Pleistocene deposits; Barrier, 1987). The Plio-Pleistocene succession starts with lower Plio- cene deep-water whitish foraminiferal marls, marly limestones, and coarser sediments including breccias, overlaid with middle Pliocene to middle Pleistocene sediments, usually in thin discontinuous, often laterally heteropic bodies. Those bodies can be grouped in: (1) a middle Pliocene–middle Pleistocene “Bathyal Facies Association” (PP), and (2) a middle Pleistocene “Circalittoral-Infralittoral Facies Association” (mP) (Barrier, 1987; Barrier et al., 1987a;Vertino,2003). PP includes carbonate- dominated and siliciclastic-dominated facies. The former facies mainly consist of coral-rich rudstones, with the frame-building deep-water scleractinians Madrepora oculata Linnaeus, 1758, Figure 1. Location of (1) Sicily in the Mediterranean Sea and (2) the study area Desmophyllum pertusum (Linnaeus, 1758), and D. dianthus in northeastern Sicily with sampling localities (Capo Milazzo, Scoppo, and the Apollo Bank, see asterisks); (3) shows Cala Sant’Antonino and Punta Mazza (Esper, 1794), interfingered with calcarenites and carbonate sections at Capo Milazzo. Modified from Rosso and Sciuto (2019). sands containing scattered isidid octocorals, and locally truncated by erosional surfaces and overlaid with debris-flow deposits. The siliciclastic-dominated facies are mainly characterized by marly Vertino, 2003). At Scoppo, these sediments unconformably lie and silty clays, sometimes embedding coral rudstone boulders on Messinian brecciated evaporitic limestone. They consist of that are often encrusted by corals, bivalves, serpulids, and bryozo- basal rudstones rich in fragments of cold-water corals (i.e., M. ans (Barrier, 1986, 1987; Barrier et al., 1996). Facies mP includes oculata, D. pertusum, and D. dianthus) that are overlain by the “upper gravels and sands” with fossils of infralittoral–upper poorly cemented white marls with sparse corals and plates of circalittoral origin and, locally, large blocks encrusted by circalit- the cirriped Scillaelepas Seguenza, 1876. These macrofossils, toral organisms, and Gilbert-type delta deposits regionally known and ostracodes, point to deposition in bathyal environments as the “Messina Formation.” The succession is erosionally capped (Vertino et al., 2013; Sciuto, 2016) in the MNN19b–19c bio- by Upper Pleistocene fluvio-marine terraces. zones (A. Baldanza, personal communication, 2015), corre- At Capo Milazzo, the so-called “yellow calcareous marl” sponding to the early Calabrian (=Santernian). crops out along the south-western and the north-eastern coast. The sandy-silty sediments unconformably lie on erosive surfaces Materials and methods of the pre-Messinian basement (Paleozoic metamorphites to upper Miocene shallow-water deposits), constituting discontinuous Studied material originates from deep-water sediments cropping sedimentary bodies filling small depressions (Fois, 1990). Sedi- out in two different localities near Messina in north-eastern ment deposition, previously dated as late Pliocene, occurred during Sicily: Capo Milazzo Peninsula (two outcrops: Cala Sant’Anto- the MPl5 and MPl6 zones, largely overlapping with the Gelasian nino and Punta Mazza) and Scoppo (Fig. 1; see Geological set- Stage of Rio et al. (1994), and now considered as the basal part ting for details). Additional material used for comparison of the Pleistocene (Gibbard et al., 2010; Violanti, 2012). Depos- derives from a present-day submarine sample collected at the ition in epibathyal environments is indicated by both macrofaunal Apollo Bank off Ustica Island in the Tyrrhenian Sea (Fig. 1). associations, including brachiopods, corals, serpulids, and, occa- At Capo Milazzo, cribrilinid bryozoans were found in sionally, mollusks (e.g., Gaetani and Saccà, 1984;Langer,1989), “sample 1 (1999)” collected near the top of the layers exposed as well as microfaunas, including foraminiferans and ostracodes at Cala Sant’Antonino West; “sample 17 (2000)” and “sample (e.g., Violanti, 1988; Sciuto, 2014a, b). Bryozoans are common, 2015” collected in the central part of Cala Sant’Antonino out- but hardly detectable in the field owing to the small size of their col- crop; and “sample 4” and “sample 5” collected in biogenic onies and/or colony fragments. Bryozoan assemblages are very layers near the base of Punta Mazza section, corresponding to diverse, including up to 60 species, some exclusively found in “sample 12” and “sample 11” of Sciuto (2014b), respectively. these deposits (Rosso, 2002a, b, 2005; Rosso and Braga, 2013; Further information on these samples can be found in Sciuto Rosso and Di Martino, 2015; Rosso and Sciuto, 2019). (2014b) and Rosso and Sciuto (2019). At Scoppo, cribrilinids Scoppo is located immediately west of the city of Messina, were found in a test sample associated with a Scillaelepas-rich in the Messina Strait area, where Pleistocene bathyal sediments layer, and in the sample “Scoppo 24 top” coming from unce- discontinuously occur (Barrier, 1984; Barrier et al., 1987a; mented marly sediment.

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At the Apollo Bank, coarse sediments associated with the Cribrilaria profunda new species kelp Laminaria rodriguezii Bornet, 1888 were collected at Figures 2, 3; Table 1 about 60 m depth. Living and dead bryozoan associations were characterized by high species richness, but delivered 1988 Puellina (Cribrilaria) scripta; Harmelin and Aristegui, only one colony (now fragmented) of Figularia figularis (John- p. 526, figs. 18–19, 24. ston, 1847) (Di Geronimo et al., 1990). 1993 Puellina scripta; Harmelin and d’Hondt, fig. 5. Sediment was routinely treated (washed, sieved, and dried) at the Paleoecological Laboratory of the University of Catania. Holotype.—PMC. B27.10.10.2019a. Capo Milazzo Peninsula: All bryozoans were picked from residues larger than 0.5 mm. Cala Sant’Antonino center, sample 2015: one small fragment After preliminary identification under a stereomicroscope, including ovicellate zooids and interzooidal avicularia. selected uncoated specimens were mounted for scanning electron microscopy (SEM) using a TESCAN VEGA 2 LMU in backscattered-electron/low-vacuum mode at the Microscopical Paratypes.—PMC. B27.10.10.2019b. Additional specimens Laboratory of the University of Catania. For the attribution of from Capo Milazzo Peninsula: Cala Sant’Antonino West, the specimens to the genera Cribrilaria Canu and Bassler, sample 1 (1999: surface): one specimen; Cala Sant’Antonino 1929 and Glabrilaria Bishop and Househam, 1987, we followed center, sample 17 (2000): three specimens; sample 2015: 12 the diagnoses in Rosso et al. (2018) summarized herein: Cribri- specimens in addition to the holotype. PMC. B27.10.10.2019c. laria has totally calcified non-pseudoporous ooecia produced by Scoppo: sample 24 top: two specimens. the distal autozooid or kenozooid, interzooidal avicularia of variable size and shape, usually five (4–8) oral spines, and rela- Diagnosis.—Colonies encrusting, multiserial. Autozooids tively large uncalcified windows of pore-chambers; Glabrilaria nearly flat, oval to irregularly polygonal. Basal pore-chambers has non-pseudoporous ooecia that are exclusively produced by present. Gymnocyst visible along the zooidal margins. Frontal the distal kenozooid, erect or semi-erect avicularia, 6–7 (rarely shield consisting of 14–25 costae with 4–11 intercostal pores/ five) oral spines, small to moderately sized uncalcified windows lacunae. Suboral bar formed by the first pair of widest costae of pore-chambers. Measurements were obtained from SEM with blunt median prominence and proximal pore. Orifice images using the image processing program ImageJ (Schneider transversely D-shaped with five (occasionally 6–7) oral spines, et al., 2012). Measurements were tabulated and provided in four in ovicellate zooids. Interzooidal avicularia with elongate, micrometers. The complete range is given first, followed by triangular or parallel-sided, raised rostrum, crossbar lacking. the mean value plus/minus standard deviation and the number Ovicell hyperstomial, presumably cleithral. Ooecium formed by of measurements taken. In specimens of Glabrilaria, zooidal distal autozooid, with a longitudinal median carina. Kenozooids boundaries were obliterated by recrystallisation with bands of rare. crystals filling the interzooidal grooves. To estimate zooidal fi size, length was measured from the distal end of the ori ce to Occurrence.—Cribrilaria profunda n. sp. is presently known the mid-point of the crystal band located proximally, while from the early Pleistocene deep-water deposits of southern width was measured from mid-point to mid-point of the crystal Italy (Gelasian of Capo Milazzo Peninsula and early Calabrian bands located laterally. of Scoppo, Messina), in the Recent Ibero-Moroccan Gulf (223–990 m depth), the Gibraltar Strait (580 m depth) Repositories and institutional abbreviations.—All specimens (Harmelin and Aristegui, 1988), and in the Alboran Sea (205 described and illustrated in this work are part of the Rosso m) (Harmelin and d’Hondt, 1992, 1993). Collection deposited at the Museum of Paleontology of the University of Catania (PMC) under the catalogue numbers reported in the “Systematic paleontology” section. Other Description.—Colonies encrusting, multiserial, unilaminar, the abbreviations: MNHN, Muséum national d’Histoire naturelle, largest observed fragment including a dozen zooids. Zooids Paris; NHMUK, Natural History Museum, London; NMNH, large and nearly flat, slightly longer than wide (L/W = 1.15: National Museum of Natural History, Smithsonian Institution, Scoppo; 1.29: Milazzo), oval to rhomboidal or rarely Washington DC. irregularly polygonal in shape, wider in their proximal half; zooidal boundaries marked by shallow grooves (Figs. 2.1, 2.4, Systematic paleontology 2.7, 3.1, 3.6). Gymnocyst exposed all along the zooidal margins, usually wider laterally to the orifice and at triple Phylum Bryozoa Ehrenberg, 1831 zooid junctions (Figs. 2.1, 3.1, 3.2). Interzooidal Order Cheilostomatida Busk, 1852 communication through basal pore-chambers with windows Suborder Flustrina Smitt, 1868 (∼70 × 20 μm), visible only in some zooids at colony Superfamily Cribrilinoidea Hincks, 1879 periphery (Fig. 3.5). Frontal shield flat (Figs. 2.1, 2.7–2.9, 3.1, Family Cribrilinidae Hincks, 1879 3.2), consisting of 14–25 wedge-shaped costae (including Genus Cribrilaria Canu and Bassler, 1929 suboral), narrowing and tapering towards the center of the zooid (maximum basal width 32–65 μm), converging toward a Type species.—Eschara radiata Moll, 1803, by original median point or along a median longitudinal, transverse, or designation. trifurcate midline. Costae connected by several intercostal

Figure 2. Cribrilaria profunda n. sp., Capo Milazzo, Gelasian. (1–4): PMC. B27.10.10.2019a, holotype with slightly recrystallized zooids, Cala Sant’Antonino center, sample 2015: (1) group of autozooids, some ovicellate, and interzooidal avicularia (ooecium shows no median carina); (2) distal part of an ovicellate zooid with four spine bases situated laterally to the orifice, the suboral bar, and intercostal lacunae; (3) close-up of autozooidal orifice with five spine bases; (4) close-up of an avicularium and a kenozooid. (5–9) PMC. B27.10.10.2019b, same details as the holotype; one of the largest paratype specimens: (5) general view (note different zooidal shapes); (6) ovicellate zooid tilted to show the median carina of the ooecium; (7) autozooids and an avicularium; (8) an autozooid with recrystallized hidden margins; (9) autozooids of different shapes. Scale bars: (1) 500 μm; (2, 3) 100 μm; (4, 6–9) 200 μm; (5) 1 mm.

bridges leaving 4–11, regularly spaced, subrectangular lacunae, between the costate shield of adjacent autozooids, no crossbar 8–16 μm long; peripheral pores the largest. Intercostal pores (Figs. 2.1, 2.4, 2.7, 2.9, 3.5, 3.6). Ovicell hyperstomial, reduced to 4–5 proximally to the first suboral pair of costae presumably cleithral. Ooecium formed by the distal autozooid. (Fig. 2.2). These are shorter and larger than the other pairs, Ectooecium smooth, with a longitudinal median elevated and merge along the zooidal midline leaving a suture with a carina (Figs. 2.1, 2.2, 2.6, 3.6). A single kenozooid with median pore, and often forming a more or less elevated costate frontal shield numbering 13 costae was observed prominence distally, adjacent to the pore (Figs. 2.2, 3.3, 3.4). (Fig. 2.4). Ancestrula not seen. Orifice transversely D-shaped, outlined by a raised rim. Orifice bearing five (occasionally up to 7) equally spaced, Etymology.—From the Latin profundus, alluding to its articulated oral spines (Figs. 2.3, 3.3, 3.4), four persisting in deep-water distribution. ovicellate zooids (Fig. 2.2). Interzooidal avicularia common, directed laterally or rarely distolaterally, with a variably Remarks.—Specimens from Capo Milazzo and Scoppo are very shaped (often triangular) cystid and an elongate triangular to similar in general appearance, including the occurrence of some almost parallel-sided rostrum, raised above or positioned irregularly polygonal autozooids with a somewhat trifurcate

Figure 3. Cribrilaria profunda n. sp., Scoppo, sample 24 top, early Calabrian, MNN19b-19c biozones, PMC. B27.10.10.2019c, paratype. (1) The largest fragment; (2) general view of an autozooid; (3) close-up of an orifice with unusual L/W ratio and seven oral spine bases; (4) orifice with five oral spine bases; (5) colony margin showing basal pore-chambers and interzooidal avicularium; (6) ovicellate zooid, avicularium, and ooecium showing longitudinal carina. Scale bars: (1) 500 μm; (2, 6) 200 μm; (3–5) 100 μm.

Table 1. Measurements (in μm) of Cribrilaria profunda n. sp. Abbreviations: L: length; W: width.

Species Cribrilaria profunda n. sp. Locality Capo Milazzo Scoppo Number of costae 17–25; 21 ± 2 (N = 14) 14–19; 17 ± 1 (N = 12) Zooid length 422–750; 640 ± 85 (N = 14) 491–711; 622 ± 64 (N = 12) Zooid width 271–676; 495 ± 108 (N = 14) 447–735; 522 ± 85 (N = 12) L/W 1.29 1.19 Proximal gymnocyst length 60–189; 96 ± 32 (N = 14) 70–246; 112 ± 50 (N = 10) Costate shield length 299–640; 407 ± 79 (N = 14) 294–442; 358 ± 45 (N = 12) Costate shield width 300–584; 450 ± 71 (N = 14) 364–586; 423 ± 61 (N = 12) Orifice length 92–97; 95 ± 4 (N = 2) 73–95; 81 ± 8 (N = 8) Orifice width 130–144; 137 ± 10 (N = 2) 115–134; 126 ± 7 (N = 8) Number of articulated oral spines 5 (4 if ovicellate) 5–7 (4 if ovicellate) Ooecium length 225–285; 261 ± 27 (N = 4) 235–253; 244 ± 24 (N = 2) Ooecium width 252–303; 271 ± 24 (N = 4) 286–320; 303 ± 24 (N = 2) Ovicellate orifice length 81–94; 86 ± 7 (N = 3) 93 Ovicellate orifice width 138–148; 142 ± 5 (N = 3) 128 Interzooidal avicularium rostrum length 225–333; 263 ± 35 (N = 12) 135–180; 158 ± 32 (N = 2) Interzooidal avicularium rostrum width 70–149; 109 ± 20 (N = 12) 57–64; 61 ± 5 (N = 2) Interzooidal avicularium cystid length 261–396; 336 ± 69 (N = 3) 273–323; 304 ± 27 (N = 3) Interzooidal avicularium cystid width 236–406; 295 ± 95 (N = 3) 154–386; 238 ± 129 (N = 3) Kenozooid length 346 Not observed Kenozooid width 339 Not observed

suture in the costate shield. Measurements also largely overlap, measurements, the presence of generally five oral spines, although Capo Milazzo material shows more variability. Yet, and presence of a robust and smooth pair of suboral costae some specimens from Scoppo show a slightly convex costate forming a median prominence. shield with fewer costae, a more raised suboral prominence, In addition, specimens from the early Messinian of Car- and more (occasionally 6–7) oral spine bases. Variability in boneras (SE Spain) identified by J.-G. Harmelin as Puellina the number of oral spines within the same species is known in (Cribrilaria) scripta and mentioned in Barrier et al. (1992), other cribrilinids, such as Cribrilaria pseudoradiata Harmelin without description or illustrations, might belong to C. pro- and Aristegui, 1988. Specimens reported as Cribrilaria funda n. sp. scripta (Reuss, 1848) by Harmelin and Aristegui (1988) and The Recent Cribrilaria pseudoradiata from the upper Harmelin and d’Hondt (1993) share their characters with the bathyal Atlanto-Mediterranean region is also similar to C. pro- Capo Milazzo material and are here considered conspecific funda n. sp., but has smaller dimensions and lacks interzooidal (see below). avicularia. Cribrilaria profunda n. sp. is very similar to the Recent C. Cribrilaria profunda n. sp. could possibly correspond to saginata Winston, 2005 from off Bahia Honda (Cuba) (Winston, Lepralia planicosta Seguenza, 1880, a cribrimorph species 2005) and the Bahama Bank (Rosso et al., 2018). However, C. reported from Plio-Pleistocene sediments of the Messina saginata differs in having a distinctly more extensive proximal Strait area. Seguenza (1880) distinguished his species from gymnocyst, a shorter and squatter orifice (orifice length/orifice C. scripta, adducing that autozooids were irregularly shaped, width 0.42–0.55 in C. saginata vs. 0.64–0.69 in C. profunda with a flat costate shield consisting of several costae, as in n. sp.), five constant oral spines, and carinated suboral costae. C. profunda n. sp. Unfortunately, Lepralia planicosta, sup- Hincks (1884), and later Neviani (1900), also suggested conspe- posedly corresponding to Lepralia scripta sensu Manzoni cificity between C. saginata,asC. radiata (Moll, 1803)from (1875) from the early Pliocene of Castrocaro, was not fig- Florida, and the middle Miocene (Langhian) Lepralia elegantis- ured and the type material was lost in 1908 during the Mes- sima Seguenza, 1880 from southern Calabria (Italy), which is, sina earthquake. We refrain from selecting our material as however, extremely unlikely owing to the great geographic the neotype of L. planicosta because the original description and temporal distance between the two populations. In addition, of this species seems insufficient to ensure their conspecifi- the only illustration available for L. elegantissima (Seguenza, city, and the type localities, although geographically close, 1880, pl. 8, fig. 11) is a drawing showing a very distinctive are not exactly the same, and neither are the geologic hori- morphology for this species, with ovoidal zooids having a zons. Seguenza (1880) abstained from illustrating his new wide and prominent frontal median keel, and seemingly 3–5 species and referred to drawings of L. scripta sensu Manzoni suboral tubercles alternating with lacunae. (1875, figs. 25, 25a). Manzoni’s specimens, held in the col- Cribrilaria scripta and C. radiata, although similar in lection of the Museo di Storia Naturale, Geologia e Paleon- appearance to C. profunda n. sp., have smaller zooidal dimen- tologia of Florence, should be located and examined before sions and larger interzooidal avicularia, and four oral spines selecting a neotype for this species. occur in most zooids in the latter species (Harmelin, 1970; Bishop and Househam, 1987). Recent specimens of C. Genus Glabrilaria Bishop and Househam, 1987 scripta sensu Harmelin and Aristegui (1988) from deep waters of the Ibero-Moroccan Bay and Gibraltar Strait, are Type species.—Puellina pedunculata Gautier, 1956, by original here attributed to C. profunda n. sp. based on the designation.

Figure 4. Glabrilaria cf. G. pedunculata Gautier, 1956, Capo Milazzo, Gelasian, Rosso Collection collective code PMC I. Pl. B.81a. (1–5) Cala Sant’Antonino center, sample 2015: (1) small fertile colony, with autozooids radiating from an apparent central ancestrula, seemingly regenerated as a miniature autozooid; (2) close-up of the three zooids on the top left of (1); note the carinate ooecia; (3) frontal view of autozooid with the transversely D-shaped oriﬁce, seven oral spines, and a recrystallized suboral area; (4, 5) inclined views of an ovicellate zooid with four oral spines and ooecium formed by the distal kenozooid with small costal shield; arrows indicate the basal pore chambers potentially producing the avicularia lateral to the ovicell; (6) Cala Sant’Antonino center, sample 17 (2000), part of a large worn colony on a bioclast; abundant kenozooids with eight costae are seen between autozooids. Scale bars: (1, 2, 6) 200 μm; (3–5) 100 μm.

Glabrilaria cf. G. pedunculata (Gautier, 1956) cf. 1966 Colletosia pedunculata; Prenant and Bobin, p. 596, Figure 4; Table 2 fig. 207 III. cf. 1970 Cribrilaria pedunculata; Harmelin, p. 93, fig. lg, h, cf. 1956 Puellina pedunculata Gautier, p. 203, fig. 20. pl. 2, fig. 6.

Table 2. Measurements (in μm) of Glabrilaria cf. G. pedunculata Gautier, 1956 and Glabrilaria transversocarinata n. sp. L: length; W: width.

Species Glabrilaria cf. G.pedunculata (Gautier, 1956) Glabrilaria transversocarinata n. sp. Locality Capo Milazzo Scoppo Number of costae 13–17; 15 ± 1 (N = 11) 14–16; 15 ± 1 (N = 10) Zooid length 252–425; 337 ± 67 (N = 10) 407–457; 436 ± 23 (N = 5) Zooid width 211–323; 263 ± 37 (N = 10) 271–337; 302 ± 27 (N = 5) L/W 1.28 1.44 Proximal gymnocyst length narrow and sloping narrow and sloping, proximal tip Costate shield length 166–257; 201 ± 28 (N = 10) 227–268; 250 ± 19 (N = 5) Costate shield width 192–277; 238 ± 31 (N = 10) 244–264; 254 ± 9 (N = 5) Orifice length 46–56; 50 ± 3 (N = 7) 45–70; 59 ± 9 (N = 4) Orifice width 69–79; 74 ± 4 (N = 7) 63–99; 83 ± 12 (N = 4) Number of articulated oral spines 7 (4 on ovicellate ones) 6 (4 on ovicellate ones) Ooecium length 134–148; 139 ± 8 (N = 3) 139–165; 151 ± 11 (N = 4) Ooecium width 159–185; 170 ± 13 (N = 3) 153–240; 194 ± 19 (N = 4) Ooecium length with kenozooid 197 186–213; 200 ± 19 (N = 2) Ooecium width with kenozooid 195 199–265; 232 ± 47 (N = 2) Ovicellate orifice length 44–47; 46 ± 2 (N = 2) 60 Ovicellate orifice width 80–86; 83 ± 4 (N = 2) 74 Kenozooid length 108–173; 119 ± 28 (N = 4) 92 Kenozooid width 94–144; 103 ± 24 (N = 4) 78

cf. 1987 Puellina (Glabrilaria) pedunculata; Bishop and autozooids, seemingly polygonal, with boundaries obliterated Househam, figs. 95–97, tab. 13. by recrystallisation, with extensive gymnocyst and costate cf. 1988 Puellina (Glabrilaria) pedunculata; Harmelin, frontal shield of 6–8costae(Fig. 4.6). The only ancestrula p. 31, figs. 9–11. found seemingly regenerated as a miniature autozooid (Fig. 4.1). cf. 2013a Puellina (Glabrilaria) pedunculata; Rosso et al., tab. 17.1. Materials.—Rosso-Collection, collective code: PMC I. Pl. cf. 2015 Puellina (Glabrilaria) pedunculata; Sanfilippo B.81a: Capo Milazzo Peninsula: Cala Sant’Antonino center: et al., tab. 2, fig. 5f. sample 2015: three specimens; sample 17 (2000): one cf. 2019a Glabrilaria pedunculata; Rosso et al., fig. 5e, f. specimen; Punta Mazza: sample 4: two specimens; sample 5: one specimen. Holotype.—MNHN-IB-2008-10384, Grand Conclu de Riou (Golfe de Marseille), Mediterranean, Recent. Remarks.—The available specimens are worn and recrystallized, preventing recognition of some diagnostic characters. However, Occurrence.—Glabrilaria pedunculata is an endemic the morphology and morphometrics of autozooids, ooecia, and Mediterranean species, widespread throughout the basin, from the kenozooids are closely reminiscent of Glabrilaria pedunculata Gulf of Lion to the Aegean Sea. Its presence in the Atlantic is Gautier, 1956, although with a few small differences. The restricted to areas swept by Mediterranean outflow water present-day Mediterranean species invariably shows six oral spines (Harmelin and d’Hondt, 1992). It has been reported from: (1) and two median pores in the triangular shelf distal to the suboral shallow-water submarine caves in the Provençal area (Harmelin, costae (Bishop and Househam, 1987, fig. 97; Harmelin, 1988, fig. 1969, 1970, 1988, 2003),intheIoniansea(Rossoetal.,2013a, b; 17a, c; Rosso et al., 2019a, fig. 5e, f). However, both the Sanfilippo et al., 2015) and Aegean sea (Crete: Harmelin, 1988; variability in the number of oral spines and the presence/absence Lesvos: Rosso et al., 2019a); (2) cryptic microhabitats from of median pores are considered to be in the range of intraspecific shallow waters (Harmelin, 2003), mid-shelf сoralligenous cliffs, variability in cribrilinids (e.g., C. pseudoradiata Harmelin and and outer shelf “Coralligène de Plateau,” at 100–140 m depth off Aristegui, 1988 and G. orientalis Harmelin, 1988). The long- Lybia and near Santorini (Harmelin, 1988); and (3) at bathyal stalked (=pedunculate) avicularia, originating from basal pore depths, ∼700 m in the Sicily Strait (Harmelin, 1979, 1988), ∼280 chambers in both autozooids and kenozooids, which are typical of m in the southern Adriatic Sea (D’Onghia et al., 2015), and ∼500 G. pedunculata, were not observed in our fossil specimens. This is m at Leuca, northeastern Ionian Sea (Mastrototaro et al., 2010), likely a taphonomic bias, because such avicularia can be easily usually associated with cold-water coral habitats. Specimens from detached even in living colonies, as observed in Glabrilaria the Gelasian of Sicily represent the first fossil record for hirsuta RossoinRossoetal.,2018 from the Bahama Bank. In our this species, suggesting its persistence, at least in deep-water fossil specimens, zooidal boundaries are mostly covered by settings, in the Mediterranean since the early Pleistocene. neomorphic calcite crystals that prevent the detection of the basal pore chambers from which the pedunculate avicularia are budded. Description.—Colony encrusting, multiserial, unilaminar However, in Figure 4.4 and 4.5 (see arrows) the pores potentially (Fig. 4.1, 4.6), the largest specimen including at least 50 producing the avicularia lateral to the ovicell are visible. zooids. Zooids oval, longer than wide (L/W = 1.28), convex, Seven oral spines were described in Glabrilaria corbula outlined by furrows filled by incipient re-crystallization Bishop and Househam, 1987 and Glabrilaria orientalis lusitanica (Fig. 4). Interzooidal communication through basal Harmelin, 1988, two closely related extant species reported from pore-chambers, more than 10 visible only in some marginal the Atlanto-Mediterranean region and the Gibraltar Strait area, zooids, with longitudinally elongate windows ∼10 × 20 μm respectively. However, the former species shows an ooecium (Fig. 4.4). Gymnocyst narrow, steeply sloping. Costate frontal that is formed by a distal kenozooid which is not distinguishable shield oval and extensive, formed by 13–17 (including in frontal view, has 4–6 costae-like ridges arranged in a radial pat- suboral) wedge-shaped, prominent costae, 27–45 μmwideat tern, a flatter autozooidal shield with somewhat carinate costae that the base, converging towards the midline and forming a slightly are sometimes with a pelma, and two large pores in the suboral raised carina (Fig. 4.4, 4.5). Costae joined by regularly spaced shelf (Bishop and Househam, 1987; Harmelin, 1988), while the intercostal bridges leaving 6–7 slit-like intercostal pores, latter species lacks midline pores in the suboral shelf (Harmelin, ∼7–8 μm long (Fig. 4.5). Only four intercostal spaces occur 1988). Glabrilaria orientalis lusitanica also has semi-erect inter- proximally to the suboral pair of costae, which are flat and zooidal avicularia (Harmelin, 1988) backed against the ooecium. merge at the midline forming a triangular shelf, possibly Six to seven oral spines also occur in Glabrilaria africana (Hay- leaving a single round pore (Fig. 4.3, 4.6). Orifice ward and Cook, 1983), but this species has numerous variably transversely D-shaped (Fig. 4.1, 4.3, 4.6), marked by a sized pores in the suboral shelf in addition to semi-erect avicularia raised rim, provided with 6–7 closely spaced, articulated oral associated with the ooecium and squeezed between autozooids. spines (Fig. 4.1, 4.3), four persisting in ovicellate zooids (Fig. 4.2, 4.5). Ovicells hyperstomial, presumably cleithral. Glabrilaria transversocarinata new species Ooecium formed by distal kenozooid, with frontally visible Figure 5; Table 2 small costate shield consisting of three costae (Fig. 4.4); ectooecium smooth, with elevated longitudinal carina Holotype.—PMC. B28.10.10.2019a: colony consisting of ∼20 (Fig. 4.2, 4.4, 4.5). Avicularia not observed. Abundant small autozooids, some ovicellate. Scoppo, sample 24 top, early kenozooids recorded in larger colonies, interspersed between Calabrian, MNN19b-19c biozones.

Figure 5. Glabrilaria transversocarinata n. sp., Scoppo, sample 24 top, early Calabrian, MNN19b-19c biozones, PMC. B28.10.10.2019a, holotype. (1) The largest specimen consisting of partly superimposed colony layers; (2) group of zooids at the colony margin showing intercostal spaces; (3) cluster of ovicellate and non-ovicellate zooids; arrow indicates a small kenozooid with five costae (note the elevated bases of oral spines and the transversely oriented crest located in the middle of the ooecium and the possible persistence of four oral spines); (4) two ovicellate zooids (note the prominent bifid suboral mucro and flat shield composed of somewhat tuberculate costae). Scale bars: (1) 500 μm; (2–4) 200 μm.

Paratype.—PMC. B28.10.10.2019b: small colony fragment and tuberculate costae with 3–7 intercostal spaces. Suboral pair of including seven autozooids, two ovicellate. Scoppo: sample costae forming a biﬁdmucro.Oriﬁce transversely D-shaped with 24 top, early Calabrian, MNN19b-19c biozones. six oral spines, four persisting in ovicellate zooids. Ovicells subimmersed. Ooecium formed by distal kenozooid, surface Diagnosis.—Colony encrusting, multiserial. Autozooids convex. smooth, with transverse rib. Avicularia not observed. Gymnocyst narrow. Frontal shield consisting of 12–14 prominent Kenozooids rare.

Occurrence.—Only known from the early Calabrian of Scoppo, (Harmelin, 1978). Oral spines are invariably seven in this Messina. species. Occasionally, transverse ornamentation has been reported Description.—Colony encrusting, multiserial, unilaminar, but in the ooecia of other cribrilinid genera. A succession of ribs including superimposed lobes (Fig. 5.1), the largest observed adds to a longitudinal carina in Puellina cassidainsis Harmelin, fragment consisting of ∼20 zooids. Zooids oval, longer than 1984 from the 3PP submarine cave in the Mediterranean French wide (L/W = 1.44), convex, the outline hidden by incipient coast (see Harmelin, 1984, fig. 7b). A cruciform pattern can recrystallization (Fig. 5.4). Interzooidal communication develop in the ooecia of Cribrilaria macaronensis (Harmelin, through basal pore-chambers visible in some peripheral 2006), and transverse ridges or wrinkles in Cribrilaria atlantis zooids, with slightly longitudinally elongate windows (Harmelin, 2006), both species previously assigned to Puellina ∼21 × 18 μm(Fig. 5.2). Gymnocyst very narrow, except for (see Harmelin, 2006, fig. 1). proximal and, occasionally, lateral extensions wedged between Measurements of Glabrilaria transversocarinata n. sp. neighboring zooids (Fig. 5.2, 5.3). Frontal shield oval and generally overlap with those of G. cf. G. pedunculata from extensive, formed by 14–16 (including suboral) wedge- Capo Milazzo (Table 2), but tend towards the higher values, shaped, prominent, tuberculate costae, 26–47 μm wide at the sometimes exceeding the upper limit. The only exception is base, converging towards the midline (Fig. 5.2–5.4). Costae the size of the kenozooid, which seems to be smaller, although joined by intercostal bridges apparently leaving 6–7 intercostal only based on a single measurement. However, morphological pores (Fig. 5.2), seemingly reduced to 3–4 proximally to the differences, including the number of oral spines, shape of costae, suboral pair of costae. These are shorter and more robust than suboral lacuna and ooecia, and the rarity of kenozooids, distin- the other costae and raised at the midline, forming a bifid guish the two species. mucro (Fig. 5.2, 5.3). Orifice transversely D-shaped, provided The two colony fragments available are detached from the with six closely spaced, articulated oral spines (Fig. 5.2), four substratum, a common feature for bryozoan specimens found in persisting in ovicellate zooids (Fig. 5.3, 5.4). Ovicells the Capo Milazzo “yellow marl.” This may indicate either that subimmersed. Ooecium formed by the distal kenozooid, with the substratum was organic or that selective aragonitic dissol- frontally visible costate (4–5 costae) shield and distal band of ution took place during/before fossilization. gymnocyst (Fig. 5.3, 5.4); ooecium with prominent, transverse, straight to slightly arched rib possibly with Genus Figularia Jullien, 1886 protruding spikes (lost) (Fig. 5.3, 5.4); an additional thinner and lower longitudinal carina was observed in a single Type species.—Lepralia figularis Johnston, 1847, by original ooecium (Fig. 5.3). Avicularia not observed. Only one designation. kenozooid was observed. It was small, polygonal, with a relatively narrow gymnocyst and costate frontal shield of five Amended diagnosis.—Colony commonly encrusting, but erect, radial costae (Fig. 5.3). Ancestrula not observed. fan-shaped, or developing erect lobes in some species. Autozooids with variably developed gymnocyst, usually wider Etymology.—From the Latin transversus, meaning transversely proximally; costate shield formed by few to numerous (up placed, and carina alluding to the typical median crest of the to 30) costae, each bearing a pelma (circular to drop-shaped or ooecium. transversely elongated) varying in size and position. Orifice with well-developed poster and condyles, dimorphic and Remarks.—The co-occurrence of a prominent transverse ridge typically larger in ovicellate zooids. Oral spines absent. on the ooecium and a bifid suboral mucro is distinctive of this Avicularia, when present, vicarious, elongate, and often species. Ooecia with a transverse ridge are known in a few spatulate, with complete crossbar. Ovicells hyperstomial or species only. One is the extant Glabrilaria hirsuta Rosso in subimmersed, cleithral. Ooecium formed by the distal Rosso et al., 2018 from the Bahama Bank, in which the autozooid or kenozooid (sometimes in the same colony), ridge is, however, very arched to subtriangular and equipped bilobate, consisting of two very large, modified costae, arched with prominent spine-like processes (Rosso et al., 2018). and meeting in the midline to form a suture and/or carina; Furthermore, in G. hirsuta, the number of oral spines (six, each costa with a wide fenestra. Interzooidal communication four persisting in ovicellate zooids) occasionally increases to via mural pore chambers in the transverse walls and seven, the costae have more obvious spine-like processes at multiporous septula in the lateral walls. Ancestrula only the periphery of the frontal shield, the suboral costae form a observed in the type species, wider than autozooids, transverse spiny crest proximal to the orifice, and kenozooids subcircular, with narrow gymnocyst encircling an extensive arranged in rows or clusters are very common (Rosso et al., opesia with differentiated orifice; no spines. 2018). In the extant Glabrilaria cristata (Harmelin, 1978) from the Hyères and Meteor banks south of the Azores, the Remarks.—The finding of a new species having morphological ooecial ridge is extremely protruding and situated more skeletal features fitting into the genus Figularia Jullien, 1886, proximally towards the orifice, contributing to form a sort of but characterized by erect colony form and a very distinctive spiny collar around the orifice together with the second pair and large ooecium formed by a distal kenozooid, led to of suboral costae. These costae bear cockscomb-like spines the examination of species currently placed in this genus that are still present but smaller than those on the other pairs (Tables 3, 4).

Downloaded from https://www.cambridge.org/core. Universitetsbiblioteket i Oslo (UiO), on 18 Dec 2020 at 15:19:10, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/jpa.2020.93 https://www.cambridge.org/core/terms Downloaded from https://www.cambridge.org/core

Table 3. List of species currently belonging to the genus Figularia with description of the main skeletal morphological characters. These species conform to the diagnosis of the genus. Abbreviations: Dim Or, Dimorphic oriﬁce; Distr, Stratigraphic distribution; E, Eocene; M, Miocene; N, number; O, Oligocene; Orig, Origin; P, Pliocene; Pl, Pleistocene; R, Recent; ZL: autozooidal length; ZW: autozooidal width; Transv. = transversal; Longit. = longitudinal; Or. = oriﬁce. Symbols in the column Orig: *ooecium formed by the distal autozooid; § ooecium formed by the distal kenozooid; ? uncertain. In the columns Suture and Dim Or the asterisk indicates the occurrence of the feature. Information is mostly compiled from the original descriptions.

. Ooecium https://doi.org/10.1017/jpa.2020.93 Vicarious Species Distr Costate shield N of costae Intercostal pores Pelmata Orig Fenestrae Suture Dim Or avicularia Additional notes Figularia arnouldi P Extensive 12 numerous 1 *§ Transv. triangular *carina Spathulate Ooecium also formed by

. Buge, 1956 with bar vicarius avicularium. Universitetsbiblioteket iOslo(UiO) F. carinata R 2/3 ZL; <1/2 ZW 10–12 ﬁssure 1 (slit-like) * Transv. * * Spathulate Fenestrae undulate, slit-like in (Waters, 1887) drop-shaped to with bar Gordon (1984). Possibly elliptical two different species. F. clithridiata R 1/2 ZL; 1/2 ZW 7–10 ? 1 (oval) § Transv. Duckfeet-shaped Ooecium with peripheral

(Waters, 1887) drop-shaped semicircle of pelma al. et Rosso seemingly belonging to the distal kenozooid. F. dimorpha R 2/3 ZL; 1/2 ZW 16 numerous 1 * Transv. oval to *carina * Ogival with bar Figuerola et al., 2018 pear-shaped F. ﬁgularis ?M–R 2/3 ZL; 4/5 ZW 9–13 ∼5 1 (circular) *§ Transv. * Spathulate Specimens in Souto et al.

(Johnston, 1847) drop-shaped to with bar (2014) possibly different —

, on irregularly oval species. F. fissa R 1/3 ZL; 1/2 ZW 8–10 1 (triangular) 1 (circular) * Transv. crescentic * Spoon-like Likely a species complex (see deep Pleistocene 18 Dec2020 at15:19:10 (Hincks, 1880) with bar Harmer, 1926, figs 20–23 and Ryland and Hayward, 1992). F. fissurata R 1/2 ZL; 2/3 ZW 3–12 fissure 1 (circular) * Transv. crescentic * * Spoon-like Canu and Bassler, 1929 with bar F. haueri M Extensive 14–18 numerous Not mentioned or * Not mentioned or *carina Absent Seemingly only differing from (Reuss, 1848) visible in fig. visible in fig. F. figularis by some

morphometrics (see ‐ , subjectto theCambridgeCore termsofuse,available at Berning, 2006) cribrilinids water F. hilli R 3/4 ZL; 4/5 ZW 5–71–2 (slit-like) 1 (drop-shaped) * Transv. oval * * Absent Ooecium including a pair of (Osburn, 1950) proximo-lateral costae. F. japonica R 3/4 ZL; 4/5 ZW 11–13 1–3 (circular) 1 (drop-shaped) * 2 pairs, transv. * * Duckfeet-shaped 7–10 costae in Yang et al., Silén, 1941 triangular with bar 2018. F. mernae R 2/3 ZL; 1/2 ZW 12–18 1 (slit-like) 1 (circular) * Longit. * * Lanceolate Uttley and Bullivant, 1972 drop-shaped with bar F. pelmatifera R 3/4 ZL; 3/4 ZW 24–30 fissure + 1–2 1 (elliptical) * Longit. * * Not observed Gordon, 1984 (elliptical) drop-shaped F. philomela R Extensive 14–16 numerous Not mentioned or * Diagonal elliptical *carina Spathulate Plastic colony morphology (Busk, 1884) visible in figs. to transv. including an encrusting drop-shaped phase (var. adnata) and bilaminar erect parts. F. rhodanica M Extensive 14–20 2 Not mentioned or * Not mentioned or *carina Spoon-like Li, 1990 visible in fig. visible in fig. with bar F. speciosa R 4/5 ZL; 3/4 ZW 12–18 3 (slit-like) 1 (slit-like) * Longit. slit-like *carina * Absent (Hincks, 1881) F. spectabilis n. sp. Pl 3/4 ZL; 3/4 ZW 8–13 3-4? (subcircular) 1 (drop-shaped) § Large * * Slightly Colony erect, flabelliform, quadrangular spathulate very large ooecium. F. tenuicosta M, ?R 2/3 ZL; >1/2 ZW 19–20 1 (slit-like) 1 (slit-like) * Longit. * * Duckfeet-shaped (MacGillivray, 1895) drop-shaped F. triangula R 2/3 ZL; <1/3 ZW 12–14 1 (slit-like) Absent ? Transv. slit-like * Not observed Powell, 1967b 11 https://www.cambridge.org/core/terms Downloaded from 12 https://www.cambridge.org/core . https://doi.org/10.1017/jpa.2020.93 Table 4. List of doubtful species currently attributed to the genus Figularia. New combinations are suggested for two species, while attribution of the remaining species awaits examination of the type material. Abbreviations: Dim Or, Dimorphic orifice; Distr, Stratigraphic distribution; M, Miocene; N, number; Orig, Origin. P, Pliocene; Pl, Pleistocene; R, Recent; ZL: autozooidal length; ZW: autozooidal width. Symbols in the column Orig: *ooecium formed by the distal autozooid; § ooecium formed by the distal kenozooid; ? uncertain. In the columns Suture and Dim Or the asterisk indicates the occurrence of the feature. Information is mostly compiled from the original descriptions. Measurements provided in μm. Additional information from Duvergier (1924), Buge (1957), Grischenko et al. (2004), Winston et al. (2014), NMNH 1, and NMNH 2. . Universitetsbiblioteket iOslo(UiO) Ooecium Nof Intercostal Vicarious New Species Distr Costate shield costae pores Pelmata Orig Fenestrae Suture Dim Or avicularia combination Additional notes Figularia ampla R 2/3 ZL; 1/3 ZW 10 fissure none * Absent *carina Not mentioned Frontal shield densely Canu and Bassler, 1928 pseudoporous F. capitifera R Vestigial, 2+2 1 (elliptical) * Single, central Spathulate Vitrimurella Frontal shield and ooecium with Canu and Bassler, 1929 suboral capitifera massive pseudopores F. contraria R 2/3 ZL; 1/2 ZW 8–11 2 1 (circular) § Two pairs of small *carina * Not observed Ovicell subimmersed. Ooecium Lagaaij, 1963 membranous areas with a pair of small oval Paleontology of Journal membranous areas centrally. Two more membraneous areas are situated on the ectooecium , on laterally – – 18 Dec2020 at15:19:10 F.? crassicostulata E Extensive 16 20 3 6 Not mentioned or * Transv. crescentic Spathulate, Canu and Bassler, 1920 visible in fig. ?no bar F. duvergieri M 2/3 ZL; = ZW 14–16 4–6 Not mentioned or * Absent/not visible * Elliptical Orifice with finely denticulate Bassler, 1936 visible in fig. with bar anter. F. echinoides O Extensive 22–24 numerous 2–3 spine-like Ovicells not observed/Absent Spathulate, Brown, 1952 no bar F. jucunda R 2/3 ZL; 3/5 ZW 8–9 1 (triangular) 1 (circular) § Pseudopores and/or *carina * Not observed Ooecium with pseudopores and/or Canu and Bassler, 1929 pelmatidia pelmatidia F. kenley M 1/2 ZL; 4/5 ZW 14–16 1 (slit-like) Visible/present only ? 2 large *carina Not observed Erect bilaminar; pelma only on :1 , subjectto theCambridgeCore termsofuse,available at Brown, 1958 on suboral costae? suboral costae

F. peltata M Extensive 15–18 numerous Absent/not visible * Absent Not mentioned Flat ooecium –

(Reuss, 1874) 23 F. planicostulata M Extensive 17 several, large Absent/not visible ? Absent/not visible * Spathulate Smooth ooecium. Canu and Lecointre, 1928 F. pulcherrima R 1/2 ZL; 1/2 ZW 9–10 3–5 1 (circular) § 2 drop-shaped, basal * Not observed Ooecium with central costate area. Tilbrook et al., 2001 lateral + 2 slit-like, cf. F. tahitiensis. median F. quaylei R Extensive 10–12 fissure 2 (circular) * 4–7 Not observed Hayamiellina Costate ooecium Powell, 1967a quaylei F. rugosa M Absent/not visible ? * Lanceolate Costate ooecium (Maplestone, 1901) no bar F. ryukuensis Pl Extensive 8–10 1 (slit-like) Absent/not visible * Pseudopores Not mentioned Pseudoporous ooecium with Kataoka, 1961 ill-defined keel F. tahitiensis R 2/3 ZL; 1/2 ZW 11 numerous 1 (circular) § 2 drop-shaped, basal *Notfigured Ooecium with central costate area. (Waters, 1923) lateral + 2 slit-like, cf. F. pulcherrima. median Rosso et al.—Pleistocene deep‐water cribrilinids 13

Figure 6. Figularia ﬁgularis (Johnston, 1847), Southern Tyrrhenian Sea, Rosso collection PMC. I. Pl. B.71.b, Apollo Bank sample. (1) Small fragment consisting of ﬁve autozooids, two ovicellate, and a vicarious avicularium; left ooecium is formed by the distal autoooid, right by the distal kenozooid with frontally visible costal shield; (2) close-up of the ooecium formed by the distal kenozooid. Scale bars: (1) 500 μm; (2) 200 μm.

Figularia was introduced by Jullien (1886, p. 608) who Puysegur Bank, off the South Island of New Zealand. The fan- designated Lepralia figularis Johnston, 1847, an Atlanto- shaped colonies of the newly discovered Figularia species from Mediterranean extant species, as the type species of the genus, Capo Milazzo, although often fragmentary (Fig. 8), show a con- and included an additional fossil species Lepralia elegantissima figuration comparable to that observed in F. mernae, with basal based on the unique drawing available (Seguenza, 1880, p. 83, zooids elongated and arranged in back-to-back adjacent pairs pl. 8, fig. 11). This latter species, depicted with oral spine (Fig. 8.1, 8.2, 8.6). The lack of a costate frontal shield, with bases, is more likely to be a species of Cribrilaria (see also no obvious evidence of breakage, in several proximal/basal Remarks on Cribrilaria profunda n. sp.). Oral spines are absent zooids, suggests that simplified polymorphs, reminiscent of in the type species F. figularis (see Soule et al., 1995, fig. 45C), those in Corbulipora MacGillivray, 1895 (see Bock and Cook, as well as in all living and fossil specimens found to date (e.g., 2001) may occur. However, the raising of the erect fan-shaped Figs. 6, 7). The absence of oral spines has also been reported portions from an encrusting phase is doubtful until encrusting almost consistently in the diagnosis of the genus, with only a colonies, or at least isolated encrusting zooids, are found. few exceptions (e.g., Gordon, 1984). Further diagnostic charac- The ooecium in Figularia is generally described as ters include a complete crossbar in the vicarious avicularia, and bivalved/bifenestrate (Ostrovsky, 2013). In F. figularis,the the presence of large, symmetrical ectooecial fenestrae and a prominent bilobate ooecium is formed by the distal autozooid, median carina in the ooecium (see Soule et al., 1995; Hayward with two costae meeting in the midline leaving a suture and/or and Ryland, 1998; Kukliński and Barnes, 2009; Yang et al., forming a slightly raised carina; each costa bearing a large, 2018). irregularly shaped and transversely elongate fenestra (membran- The erect colony-form has never been mentioned in the ous area in non-cleaned specimens). The colony fragment of generic diagnosis before. However, Busk (1884, p. 132) F. figularis from the Apollo Bank (Tyrrhenian Sea, Mediterra- described Figularia philomela as “free; erect or decumbent nean) shows that ooecia formed by the distal kenozooid can (hemescharan).” Subsequently, Hayward and Cook (1979, co-occur in the same colony in this species (Fig. 6). Though p. 76) found a bilaminar fragment of F. philomela interpreted uncommonly reported, and here recorded in F. figularis for as part of an erect foliaceous colony possibly arising from an the first time, the co-occurrence of ooecia produced by the distal encrusting phase (var. adnata of Busk, 1884). Gordon (1989, autozooid and kenozooid is known in other cribrilinids, such as p. 15, 16) recorded the occasional occurrence of an erect bila- Cribrilina punctata (Hassall, 1841), “Puellina” harmeri Ristedt, mellar lobe, arising from the adjacent encrusting zooids, in a col- 1985 (see also discussion in Rosso et al., 2018), Cribrilaria ony of Figularia mernae Uttley and Bullivant, 1972 from innominata (Couch, 1844) (see Chimenz Gusso et al., 2014),

Figure 7. Figularia ﬁgularis (Johnston, 1847), Scoppo, sample 24 top, early Calabrian, MNN19b-19c biozones, Rosso collection PMC I. Pl. B.71.c. (1) Fragment with few autozooids (note the teratologic autozooid); (2) close-up of the distal half of the teratologic autozooid shown in (1); (3) fragment with four, incomplete autozooids; (4) close-up of the oriﬁce. Scale bars: (1, 3) 500 μm; (2, 4) 200 μm.

Figure 8. Figularia spectabilis n. sp., Capo Milazzo, sample Cala Sant’Antonino center, 2015, Gelasian, PMC. B22. 5.4.2015.b, paratypes, colony morphology. (1, 2) Lateral view of two fan-shaped colony fragments with thin cylindrical proximal base; (3) side view of a narrow ribbon-like fragment; (4, 5) inclined proximal view and lateral view of fan-shaped colony fragments with slightly diverging sides; (6) proximal view of a fan-shaped colony fragment; (7, 8) inclined distal and top view of a fan-shaped colony fragment. Scale bars: (1, 7, 8) 1 mm; (2–6) 500 μm.

Puellina saldanhai Harmelin, 2001, and Puellina mikelae Har- is situated in the proximal part of the distal zooid predominantly melin, 2006. Following Rosso et al. (2018), the latter two spe- below the colony surface, thus corresponding to endozooidal cies are here allocated to the genus Cribrilaria: Cribrilaria type. Whether this position of the brood cavity was an effect saldanhai (Harmelin, 2001) n. comb. and Cribrilaria mikelae of decalcification of the skeleton (and, thus, sagging of the ori- (Harmelin, 2006) n. comb. Both ovicell variants sometimes ginally raised ooecium) during preparation for sectioning is cur- may appear within the same colony (e.g., in C. punctata and rently not clear, but this contradicts most descriptions showing “P.” harmeri) indicating a developmental plasticity of this char- hyperstomial ovicells in this species (see references above). acter (reviewed in Ostrovsky, 2013). A similar plasticity in ovi- Still, a degree of the brood cavity immersion may vary, and, cell formation is only known in some Calloporidae (Ostrovsky for example, both hyperstomial and subimmersed ovicells are and Schäfer, 2003; Ostrovsky et al., 2009; Ostrovsky, 2013) known within the genus Figularia, and hyperstomial, subim- that are presumed ancestors of cribrilinids. mersed, and endozooidal ovicells are described in the different The kenozooid producing the ooecium in F. figularis shows species of Puellina (Ostrovsky, 2013). Subimmersed ovicells a crescent-shaped shield of short radial costae, each with a single were present in Recent colonies of F. figularis from the Mediter- pelma as in the autozooids, but also with a single intercostal pore ranean (A. Ostrovsky, personal observations). (Fig. 6). The same structure is also evident in the fossil species Ostrovsky and Taylor (2005) noted the occurrence of spe- from Capo Milazzo (Fig. 9). Ovicells with ooecia formed by the cies of Figularia—F. clithridiata (Waters, 1887), F. tahitiensis distal kenozooid also occur in other species currently assigned to Waters, 1923, and F. pulcherrima Tilbrook, Hayward and Gor- this genus, based on examination of available SEM images and, don, 2001—having costate ooecia (see also Ostrovsky, 2002). to a lesser extent, drawings (see Table 3). Winston et al. (2014) remarked that the occurrence of costate Ostrovsky (2013, fig. 1.28A) illustrated sectioned decalci- ooecia in F. pulcherrima possibly suggests a better allocation fied ovicells of F. figularis in which most of the brood cavity of this species in a distinct genus. Inclusion of costae in the

Figure 9. Figularia spectabilis n. sp., Capo Milazzo, sample Cala Sant’Antonino center, 2015, Gelasian, PMC. B22. 5.4.2015.a, holotype, ooecium. (1) Colony fragment with unique ovicellate zooid and vicarious avicularium; (2) close-up of the ovicellate zooid with ooecium formed by the distal kenozooid. Scale bars: (1) 500 μm; (2) 200 μm.

construction of the ooecium has also been observed in Figularia ancestrulae in additional species is needed to confirm whether hilli (Osburn, 1950), with two small costae similar to those of this morphology is constant among congeners, which has been the frontal shield added proximally to the larger ooecial halves proven not to be the case in other cheilostome genera, such as (see Table 3). e.g., Escharina Milne Edwards, 1836 (see Berning et al., 2008). Yang et al. (2018), while including pseudoporous ooecia in Several species previously assigned to Figularia were the diagnosis of Figularia, also suggested the examination of recently displaced in different genera of the families Cribrilinidae species with multiple ectooecial pseudopores in order to deter- and Calloporidae (e.g., Vitrimurella, Reginella Jullien, 1886, mine if they are genuinely congeneric. These species are here Inferusia Kukliński and Barnes, 2009, Valdemunitella Canu, re-assigned to different genera (see also below and Table 4). 1900; see Bock and Gordon, 2020), and Jullienula Bassler, Suboral costae often differ from the other pairs. In the type 1953 (Yang et al., 2018). Here, we suggest further displacements: species of Figularia, suboral costae merge, forming a smooth, both Figularia? ampla Canu and Bassler, 1928, only tentatively wide shelf facing the orifice, most evident in ovicellate zooids included in Figularia when first described, and Emballotheca? (Fig. 6). Wide suboral costae associated with ovicellate zooids capitifera, Canu and Bassler, 1929, subsequently referred to his were also observed in F. rhodanica Li, 1990.InF. pelmatifera new genus Calyptotheca by Harmer (1957)andtoFigularia by Gordon, 1984 the suboral pair of costae develops into two lateral, Di Martino and Taylor (2018), fitbetterinVitrimurella,owing divergent, spinose processes (see Gordon, 1984, pl. 19, fig. E). to the pseudoporous zooidal gymnocyst and ooecia, and the A certain variability occurs in the presence/absence of pel- extremely reduced costate shield. Figularia ryukyuensis Kataoka, mata in the frontal shield, and in their position along the costal 1961 and F. jucunda Canu and Bassler, 1929 also need to be length. Sometimes this variability was noted (e.g., Gordon, revised, pending examination of the type material. These species 1984). Nevertheless, all Figularia species lacking pelmata (i.e., have pseudoporous ooecia formed by the distal kenozooid with- not included in formal descriptions and/or undetectable in avail- out a visible frontal part. Figularia duvergieri Bassler, 1936 has able images) are fossil, except “F. philomela var. adnata” (Busk, an unusual denticulate proximal orifice margin, and lacks costal 1884), suggesting that their absence may be a preservation artefact. pelmata and fenestrae in the ooecium. A detailed revision based The ancestrula is generally not mentioned in species on SEM images is needed to confirm generic allocation for descriptions to our knowledge. In the amended diagnosis, we these problematic species (Table 4). This issue has been partially include characters of the ancestrula for the first time, based on addressed by López Gappa et al. (in press). the ancestrula found in a colony of F. figularis from the Mediter- ranean illustrated in Rosso et al. (2019b, fig. 5C). The large size Figularia figularis (Johnston, 1847) of both autozooids and ancestrula (0.65 × 0.67 mm) and the Figures 6, 7; Table 5 absence of spines are rare and remarkable among cribrilinids, which usually have small, tatiform ancestrulae, and this may 1847 Lepralia figularis Johnston, p. 314. have implications on the systematics/phylogeny of this genus 1966 Figularia figularis; Prenant and Bobin, p. 604, fig. 2010 within the family Cribrilinidae. However, observation of I–IV, VI.

Table 5. Measurements (in μm) of Figularia ﬁgularis and Figularia spectabilis n. sp. *Refers to an aberrant zooid (see text for further explanation). L: length; W: width.

Species Figularia figularis (Johnston, 1847) Figularia spectabilis n. sp. Locality Scoppo Capo Milazzo Number of costae 10–20; 14 ± 5 (N = 3)* 7–14; 10 ± 2 (N = 18) Zooid length 858 588–1057; 759 ± 135 (N = 16) Zooid width 402 319–525; 442 ± 55 (N = 16) L/W 2.13 1.72 Proximal gymnocyst length 41–111; 86 ± 31 (N = 4)* 60–210; 114 ± 47 (N = 14) Costate shield length 455–457; 456 ± 1 (N = 2) 294–582; 388 ± 93 (N = 14) Costate shield width 307–396; 352 ± 63 (N = 2) 306–543; 378 ± 65 (N = 14) Orifice length 202–228; 213 ± 12 (N = 4) 176–292; 236 ± 32 (N = 18) Orifice width 202–244; 225 ± 21 (N = 4) 179–295; 233 ± 31 (N = 18) Number of articulated oral spines absent absent Ooecium length not observed 702 Ooecium width not observed 730 Ovicellate orifice length not observed 297 Ovicellate orifice width not observed 323 Interzooidal avicularium length not observed 473–642; 566 ± 86 (N = 3) Interzooidal avicularium width not observed 273–337; 298 ± 34 (N = 3)

1998 Figularia figularis; Hayward and Ryland, p. 338, fig. Armaçao de Pêra in Portugal (Souto et al., 2014) needs to be 120, cum syn. verified. This colony has an unusual triangular ooecial 2002 Figularia figularis; Hayward and McKinney, p. 38, fig. fenestra with narrow horizontal part and could represent a 16 D–E. different species. 2006 Figularia figularis; Berning, 2006, p. 49, pl. 3, figs. 7, 10, cum syn. Figularia spectabilis new species 2014 Figularia figularis; Chimenz Gusso et al., p. 167, fig. Figures 8–11, Table 4 84a–c. Holotype.—PMC. B22. 5.4.2015.a: bilaminar fragment including Holotype.—NHMUK 1847.9.16.39, English Channel, Recent. some autozooids and the only observed ovicell. Cala Sant’Antonino, sample Cala Sant’Antonino center, 2015, Gelasian. Occurrence.—Figularia figularis is widely distributed in the Atlanto-Mediterranean area since the middle Miocene Paratypes.—PMC. B22. 5.4.2015.b: additional 39 fragments (Moissette et al., 1993; Berning, 2006). This species has from the same sample as the holotype, including several been commonly reported from shelf habitats, mostly from fan-shaped colony portions. One fragment from sample 17 the deep shelf, often associated with deep coralligenous (2000), Cala Sant’Antonino center. facies (Di Geronimo et al., 1990; Ballesteros, 2006), and at the shelf break in both the Mediterranean (110–145 Diagnosis.—Colony erect, bilaminar with fan-shaped fronds, m; see Harmelin and d’Hondt, 1992) and the eastern the tapering proximal terminations possibly consisting of Atlantic as far north as the British Isles (Hayward and heteromorphs, likely rising from an encrusting phase. Zooidal Ryland, 1998). frontal shield consisting of flat costae, each with a large, elongate drop-shaped pelma placed on its peripheral half; Materials.—Rosso collection PMC. I. H. B.71.b, Apollo Bank gymnocyst wider laterally and proximally, narrower distally, sample: two specimens, Recent; Rosso-Collection PMC I. Pl. with faint striations. Vicarious avicularia elongate, spatulate, B.71.c, Scoppo: sample 24 top: two specimens, early with extensive rostral palate and complete crossbar. Ovicell Calabrian, MNN19b-19c biozones. subimmersed, presumably cleithral. Ooecium formed by the distal kenozooid with frontally visible costate part, and Remarks.—Two fossil fragments were found, each consisting of consisting of two very large, wing-shaped costae merging a few zooids (Fig. 7). Zooidal morphological characters allow a in the midline producing a longitudinal suture, with two reliable identification, even in the absence of ovicells and large fenestrae exposing wide areas of endooecium; the avicularia. Morphometrics fall within the ranges reported for costae of the ooecium-producing kenozooid smaller, this species. Inferred teratology in an autozooid resulted in a forming a distal, crescent-shaped crown, each costa with a double-bifurcated frontal shield (Fig. 7.1, 7.2). This unusual small pelma. feature also occurs in the type specimen of F. tenuicosta MacGillivray, 1895 from the middle Miocene of Victoria, Occurrence.—Exclusively known from early Pleistocene Australia (Bock, 2020). Although F. figularis exhibits a (Gelasian) deep-water sediments of Capo Milazzo (NE Sicily, Italy). certain range of morphological variability, some historical records, mostly beyond its confirmed geographical range, Description.—Available colony fragments bilaminar, fan-shaped proved to be different species (e.g., Brown, 1952). (the largest ∼2 mm long by 3 mm wide); fragments diverging The conspecificity of the colony found on a rock at distally at variable angles from a subcylindrical proximal

Figure 10. Figularia spectabilis n. sp., Capo Milazzo sample Cala Sant’Antonino center, 2015, Gelasian, PMC. B22. 5.4.2015.b, paratypes, autozooids. (1) Frag- ment of a bilaminar branch with zooids arranged in longitudinal rows and distal vicarious avicularium; (2) group of autozooids; (3) close-up of elongated autozooid with well-defined boundaries and growth lines in the gymnocyst (note the smooth texture of the costae, converging towards the midline, and the elongate pelmata); (4) wider autozooid with large wedge-shaped costae and very large drop-shaped pelmata; (5) close-up of some costae; (6) orifice; (7) orifice with closure plate or calcified operculum. Scale bars: (1, 2) 500 μm; (3, 4) 200 μm; (5–7) 100 μm.

portion, consisting of four zooids arranged in back-to-back pairs by an uncertain number of intercostal bridges, presumably 3–4 (Fig. 8.1, 8.2, 8.4–8.8). Other fragments of similar size include (Fig. 10.5), with small oval to subcircular intercostal pores in only the edges of presumably ribbon-like colonies (Figs. 8.3, between. A longitudinal suture marking the costal fusion 10.1). Putative proximal heteromorphs, possibly arising from an along zooidal midline (Fig. 11.1). Each costa bearing a single, encrusting phase and forming the basal stalk, lacking calcified elongate, drop-shaped pelma with the rounded base placed in frontal shield. Zooidal boundaries marked by grooves. Zooids correspondence with the base of the costa, while the acute large, about twice as long as wide (L/W = 1.72), gently arched vertex extends up to half to two thirds of costal length. Orifice distally, wedged proximally. Gymnocyst more extensively oval to round, slightly longer than wide, concave proximally, exposed proximally and laterally (Figs. 8.2, 8.3, 8.5, 10.1–10.4), gently arched distally, outlined by a rim of calcification locally obliterated by recrystallization (Fig. 10.5). Costate (Fig. 10). Oral spines absent. Avicularia vicarious, infrequent, shield extensive (∼75% of the frontal surface), gently convex, elongate and slightly asymmetrical, varying in size; rostrum formed by 7–14 flat and smooth costae (maximum basal width long, spatulate, directed distally and slightly inclined, facing 72–111 μm), varying from short and subtriangular proximally frontally (Figs. 9.1, 11); post-mandibular area short, palate to long and parallel sided distally; the suboral pair often the wide, crossbar complete (Fig. 11.3). Ovicell subimmersed, largest (Fig. 10.1–10.5). Costae defined by grooves, connected presumably cleithral. A single observed ooecium formed by

Figure 11. Figularia spectabilis n. sp., Capo Milazzo, sample Cala Sant’Antonino center, 2015, Gelasian, vicarious avicularia. (1) Holotype PMC. B22. 5.4.2015.a; (2, 3) paratypes PMC. B22. 5.4.2015.b, same details as the holotype (note the spatulate rostrum and the thin crossbar); (3) view showing the wide rostral palate. Scale bars: 200 μm.

the distal kenozooid with frontally visible costate shield of 10 Cribrilinids are generally rare in Plio-Pleistocene associa- costae, longer than wide, wider and slightly more prominent tions from deep-water environments in Sicily and Calabria, as than the ovicellate zooid (Fig. 9). Very large ooecium well as in their enclaves in shallow waters, such as past submar- consisting of two flat, wing-shaped costae converging along ine cave habitats, from which a single species, Cribrilaria the midline, the fusion marked by a longitudinal suture, venusta (Canu and Bassler, 1925), and undetermined cribrilinid distally with two small tubercle-like prominences. Large taxa were previously reported (Di Geronimo et al., 1997, 2005; rhomboidal fenestra exposing finely granular endooecium. Rosso, 2005; Rosso et al., 2015). Thus, this study raises the total Orifice of the ovicellate zooid slightly larger than those of number of cribrilinids from these paleoenvironments to six spe- autozooids, rounded rectangular. Closure plates or calcified cies in three genera. Shallower shelf paleoenvironments from the opercula sometimes occluding orifices (Fig. 10.7). same regions, mostly Pleistocene but as old as Miocene, yielded seven species of cribrilinids: Cribrilaria radiata (Moll, 1803), Etymology.—From the Latin spectabilis, meaning remarkable, C. hincksi (Friedl, 1917), C. innominata (Couch, 1844), Puel- exceptional, alluding to the distinctive architecture of the lina gattyae (Landsborough, 1852), Distansescharella seguen- colony and ooecium. zai Cipolla, 1921, Gephyrotes moissettei Di Martino and Rosso, 2015, and “Cribrilina punctata” (Hassall, 1841), the lat- Remarks.—The morphology of the colony, zooids and ter species probably being a Collarina (Barrier et al., 1987b; ooecium distinguish Figularia spectabilis n. sp. from Harmelin et al., 1989; Di Geronimo et al., 1994; Rosso and San- congeners. The flabellate to short, ribbon-like morphology of filippo, 2005; Di Martino and Rosso, 2015). As for other taxa the colony, with putative heteromorphs placed basally, may authored by Seguenza (1880), the loss of the type material suggest the occurrence of basal rhizoids for fixation to the makes it difficult to confirm the status of some cribrilinid spe- substratum. Alternatively, the connection to an encrusting cies, such as Lepralia thiara, L. mitrata, and L. mitrata portion may develop through “sites of articulation” as in v. radians, in addition to the previously mentioned L. elegantis- Bryobaculum carinatum Rosso, 2002a, occurring in the same sima and L. planicosta. Analogously, the real identity of some sediment. other species (briefly described and lacking illustrations) in Waters (1878), De Stefani (1884), Hincks (1884), and Neviani Discussion (1900, and references therein) is doubtful. Focusing only on deep-water assemblages, cribrilinids are Five species of cribrilinid bryozoans, three of which are new to present with three species in both the Gelasian associations science, namely Cribrilaria profunda n. sp., Glabrilaria cf. from Capo Milazzo and the Calabrian (MNN19b-19c biozones) G. pedunculata, G. transversocarinata n. sp., Figularia figu- of Scoppo. These figures are comparable to those found in laris, and F. spectabilis n. sp., were found in Pleistocene deep- present-day deep-water associations from the Mediterranean water sediments from north-eastern Sicily. and Atlantic (Bahama Bank), in which cribrilinids usually Figularia figularis was already recorded from the area by occur with 2–3 species (Rosso et al., 2018). However, the Seguenza (1880) and Neviani (1900), while C. profunda Gelasian of Capo Milazzo includes at least 46 cheilostome spe- n. sp. was possibly recorded as Lepralia planicosta (see cies, and the cribrilinid relative percentage is ∼6%, which is Remarks above), while the remaining three species, including lower than the 10–18% found in present-day assemblages G. cf. G. pedunculata, represent new records. (Rosso and Sciuto, 2019). No comparison can be made for

the Calabrian of Scoppo whose bryozoans are still under previous projects. E. Di Martino received funding from the investigation. European Research Council (ERC) under the European Union’s Discovery of a new species of Figularia, F. spectabilis Horizon 2020 research and innovation program (grant agree- n. sp., led to the emendation of the genus diagnosis and the ment No 724324 to L.H. Liow). A. Ostrovsky thanks the re-examination of the 32 species and one variety currently Russian Science Foundation (grant 18-14-00086) for assigned to the genus, based on drawings and photographic financial support of the studies on cheilostome brood chambers. material available from the literature. This preliminary survey This is Catania Paleontological Research Group: contribution allows us to confidently reassign two species based on published n. 465. scanning electron micrographs of the type material. The newly proposed combinations are Vitrimurella capitifera (Canu and References Bassler, 1929) n. comb. and Hayamiellina quaylei (Powell, 1967a) n. comb., as also suggested by Kukliński et al. (2015). Ballesteros, E., 2006, Mediterranean coralligenous assemblages: a synthesis of present knowledge, in Gibson, R.N., Atkinson, R.J.A., and Gordon, J.D.M., Thirteen species remain doubtful and their assignment to more eds. Oceanography and Marine Biology: An Annual Review: Boca Raton, suitable genera requires examination of the type material CRC Press and Taylor and Francis Group, v. 44, p. 123–195. (Table 4). Barrier, P., 1984, Evolution tectono-sédimentaire pliocène et pléistocène du Détroit de Messine (Italie). [Ph.D. dissertation]: Marseille, France, Univer- At present, 18 species, including Figularia spectabilis sité Aix-Marseille II, 270 p. n. sp., match the diagnosis of the genus. This figure will likely Barrier, P., 1986, Evolution paléogéographique du Dètroit de Messine au Plio- cène et au Pléistocène: Giornale di Geologia, s. 3, v. 48, p. 7–24. change further after a more detailed revision of some fossil spe- Barrier, P., 1987, Stratigraphie des dépôts pliocènes et quaternaires du Détroit cies and species left in open nomenclature (see Berning, 2006 de Messine: Documents et Travaux Institut Gèologique Albert de Lappar- for F. haueri and F. figularis; Di Martino et al., 2017 and ent, Paris, v. 11, p. 59–81. Barrier, P., Di Geronimo, I., and Montenat, C., 1987a, The Messina Strait Cook et al., 2018 for two different Figularia spp.) as well as (Italy): Pliocene and Pleistocene tectono-sedimentary evolution and present cryptic species/species complexes (e.g., F. clithridiata and environment: Documents et Travaux de l’Institut Géologique Albert de Lap- F. fissa). Based on our literature review, the diversity parent, Paris, v. 11, 272 p. 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Of Barrier, P., Di Geronimo, I., La Perna, R., Rosso, A., Sanfilippo, R., and Zibro- wius, H., 1996, Taphonomy of deep-sea hard and soft bottoms the 12 species of Figularia living today, 10 species are found communities: the Pleistocene of Lazzàro (Southern Italy), in Meléndez, in the Pacific and Australasian region. Only two species, F. fig- G., Blasco, F., and Pérez, I., eds., Tafomonia y Fossilizaciòn, II Reuniòn, ularis and F. dimorpha, fall outside this area, being recorded in Saragoza: Comunicaciòn de la II Reunión de Tafonomia y Fosilizaciòn, p. 39–46. the Atlanto-Mediterranean and southwestern Atlantic regions, Bassler, R.S., 1936, Nomenclatorial notes on fossil and recent Bryozoa: Journal respectively. of the Washington Academy of Sciences, v. 26, p. 156–162. Bassler, R.S., 1953, Bryozoa, in Moore, R.C., ed., Treatise on Invertebrate Pale- A twofold future investigation is sought. This includes an ontology. 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